and to a combined reference panel including haplotypes from the 1KG Project and the UK10K Project. To improve variant coverage in non-EUR participants of UKB, we re-imputed their genotypes to the 1KG reference panel, as described previously38. Lifelines samples were imputed to the HRC panel. PAGE and CKB were imputed to the 1KG reference panel. Standard quality control (\documentclass[12pt]{minimal} \usepackage{amsmath} \usepackage{wasysym} \usepackage{amsfonts} \usepackage{amssymb} \usepackage{amsbsy} \usepackage{mathrsfs} \usepackage{upgreek} \setlength{\oddsidemargin}{-69pt} \begin{document}$${r}_{{\rm{INFO}}}^{2}$$\end{document}rINFO2 > 0.3, PHWE > 10−6 and MAC > 5) were applied to imputed genotypes in each dataset.